Botanica Marina Vol. 45, 2002, pp.


© 2002 by Walter de Gruyter · Berlin · New York

A Revision of Amansia glomerata C. Agardh, Amansia rhodantha (Harvey) J. Agardh and Melanamansia glomerata (C. Agardh) R. E. Norris (Rhodophyta: Rhodomelaceae)
A. D. R. N’Yeurt
Jeune Equipe Terre Océan, Université de Polynésie Française, BP 6570, Faa’a Aéroport, Tahiti 98702, French Polynesia,

Species of Amansia and Melanamansia from the tropical Pacific region have been re-examined, and compared with material from the Indian Ocean, Japan, Taiwan and the Philippines. All specimens examined from the Pacific region (except Hawaii and some from New Caledonia and Fiji) lack pseudo-pericentral cells, thus placing them in Amansia. Melanamansia glomerata is only reported from Hawaii, New Caledonia, Fiji, the Philippines,Vietnam, Malaysia and for the first time in the Indian Ocean from Kenya.Type material of Amansia rhodantha (Harvey) J. Agardh from Mauritius was compared and found not significantly different from the Pacific Amansia material (including type and isotype material of Amansia paloloensis South et Skelton from Samoa, which is reduced to synonymy), thus extending the distribution range of A. rhodantha as a pantropical entity.

The genus Amansia was erected by Lamouroux (1809: 322) and is placed in the tribe Amansieae of the family Rhodomelaceae (Hommersand 1963: 335). The tribe is characterised by dorsi-ventral, erect thalli arising in tufts from a common basal disc. Most genera have five pericentral cells around the axial cell (Figs 1, 2) except Kuetzingia Sonder and Protokuetzingia Falkenberg, which have six pericentral cells (Kylin 1956). The two first-formed pericentral cells lie on the dorsal side of the characteristically inrolled apex of the mature lanceolate blades. In Amansia the dorsal and dorsi-lateral pericentral cells further divide to form immediately behind the apex a distromatic wing or ala, whose cells are arranged in alternate layers, with secondary lateral pit-connections (Fritsch 1945: 570). Recently, Norris (1988b, 1995) re-examined species of Amansia in the Indian Ocean and Hawaii, and discovered that some members of this genus differed consistently by having the first and second pericentral cells each producing a lateral pseudo-pericentral cell lying adjacent, but not connected to, the axial cell (Norris 1988b: 214; Fig. 3). Species previously ascribed to Amansia that had these pseudo-pericentral cells were transferred to a new genus (Melanamansia R. E. Norris 1988b: 217). Notably, Hawaiian plants previously ascribed to the common tropical species Amansia glomerata C. Agardh (whose type locality is Hawaii) were transferred to Melanamansia glomerata (C. Agardh) R. E. Norris, while Indian Ocean ‘Amansia glomerata’ were put under the earlier name Amansia rhodantha (Harvey) J. Agardh based

on their lack of pseudo-pericentral cells (Norris 1995). The genus Amansia currently consists of four species, the Caribbean A. multifida Lamouroux (the type of the generic name), A. loriformis R. E. Norris and A. rhodantha (Harvey) J. Agardh from the Indian Ocean and Malaysia, and A. paloloensis South et Skelton from Samoa. Melanamansia currently consists of twelve species, M. daemelii (Sonder) R. E. Norris, M. glomerata (C. Agardh) R. E. Norris, M. japonica (Holmes) R. E. Norris, M. mamillaris (Lamouroux ex C. Agardh) R. E. Norris, M. pinnatifida (Harvey) R. E. Norris, M. pumila (Sonder) R. E. Norris, M. scalpellata (Tanaka) R. E. Norris, M. seagriefii R. E. Norris, M. serrata (Harvey) R. E. Norris and M. mitsuii (Tanaka) Yoshida. Following these revisions by R.E.Norris,several authors (Yoshida et al. 1995,N’Yeurt et al. 1996,Payri and N’Yeurt 1997, Millar 1999, Millar et al. 1999) made the nomenclatural change from Amansia glomerata to Melanamansia glomerata in their checklists and herbarium collections from the tropical Pacific, but until recently no detailed studies verifying these changes as correct have been done. In 1995, samples of ‘Amansia glomerata’ from Rotuma Island (north of Fiji) were sent to Dr R.E.Norris,who confirmed (pers. com.) that pseudo-pericentral cells were present, and on the basis of this assumed finding the name Melanamansia glomerata was used in later publications (N’Yeurt 1996,N’Yeurt et al. 1996).Later,in the course of on-going investigations on the Samoan algal flora, South and Skelton (1999) found a species of Amansia lacking pseudo-pericentral cells, which they described as new (A. paloloensis South et Skelton), while in Malaysia and Vietnam (Masuda et al. 2000, Masuda

3. N’Yeurt (SUVA-A 5128). 3899). a detailed morphological and taxonomic study on some selected members of the Amansieae (Wilson and Kraft 2000) echoed the opinion that the genus Amansia in Australia was in need of critical revision. P. Fiji. leg. New Caledonia. 25 October 1984 (SAP 046978). . A. Taharaa. 1997. leg. Payri (8/84. M. Results Representative material examined Amansia rhodantha – Japan: Hanasado Beach. E. Kagoshima: 9 August 1959. 13 May 1982. Punaauia. D.. SUVA-A 5438L). 19 Mar. 3895. 9 Feb. it was decided to revise records in the light of recent findings. Okidomari. 18 Sep.‘Al’ and ‘CG’ refer to specimens housed at IRD. M. 3897). Material and Methods Specimens were collected by reef-walking. 3893. and stained with cotton blue in lactophenol or 1% acidified aniline blue in 60% clear corn syrup. 5481L (spermatangial)]. cross section of the lectotype from the Hawaiian Islands (LD 42600) showing the first two dorsal pericentral cells each developing a pseudo-pericentral cell (ps) that lies adjacent to the axial cell. deep algal rim. Afiti and D. 11 Jul. Melanamansia glomerata. P. in particular paying attention to important anatomical features such as the presence or absence of pseudo-pericentral cells and the number of segments between lateral veins of the blade (Norris 1995). Rarotonga. A. Fig. – Samoa: Palolo Deep Reserve. and a camera-lucida attachment was used to make anatomical drawings.) recent studies found both Melanamansia glomerata and Amansia rhodantha (Harvey) J. Fiji. Tanegashima Island. Prepared slides were examined using a Zeiss compound microscope. 3896. R. leg. Tahiti. A. 1989. com. Skelton and Dave (SUVA-A 1567).Agardh to co-exist in the flora. 1998. A. Baba (SAP 056160). Amansia rhodantha (Beqa. N’Yeurt (UPF 347). showing intrusion of the first derivative of the first-formed pericentral cell into the axial ring (Suva. in EPHE). leg.232 A. Skelton and Dave [SUVA-A 1565 (holotype). (1981). N’Yeurt In order to clarify the taxonomy of species hitherto ascribed to Melanamansia glomerata. A. Amansia rhodantha and Amansia glomerata. 3892. The results of these investigations are described below. snorkelling or SCUBA diving. SUVA-A 1566 (isotype). otherwise as in Holmgren et al. Tahiti. Okinoerabujima Island (60 km north of Okinawa) 27 Sept. Fig. 1984. – Taiwan: Anon. Faletose (SUVA-A 3894. 5480L (cystocarpic). 2 Jul. Sakai and S. Figs 1–3. Amansia rhodantha. P. Arrangement of pericentral cells in species of Amansia and Melanamansia (modified from Norris 1988b. Sections were made using a freezing microtome. P. Most recently. leg. Rintels (SUVA-A 3891. – French Polynesia: Moorea. and stored in 5% buffered Formalin in seawater prior to shipment to the laboratory. 1. 1–5 = order of formation of pericentral cells. leg. 1998. SUVA-A 5436L). 3898. Enomoto (SAP). Skelton. 1998. 1998. Okinoerabujima Island (60 km north of Okinawa). M. Fig. leg. pers. Tombleson and T. developed and printed in the laboratory. Baba (SAP 056163). Y. leg. Skelton. leg. Nouméa. leg. ps = pseudo-pericentral cells). 6 Jun. a = axial cell. C. A. Photomicrographs were obtained using an Olympus BX-50 compound microscope. – Cook Islands: Ngatangiia. 2. 1998. 1999. U. 2 Jul. Macrophotography was done with a Nikon F2A in conjunction with Kodak Plus-X pan 125 film. leg. Palolo Deep. A. SUVA-A 2402 (Lectotype) and liquid-preserved isotypes 5479L (tetrasporic). Furusato. N’Yeurt (UPF 584).

Lectotype. 21 Sep. 1819. M. M.3 2. leg. S. Huisman (MURU. Silva et al. leg. Huisman (MURU. A. Viti Levu. South in lit. leg. B. n. – Australia: Heron Island QLD. A. Namada. Kipukai.Negros Oriental. Melanamansia glomerata – Hawaiian Islands: Hawaiian Islands. – Lord Howe Island: 18 Aug. Snellius-II (L 0108602). Unia. leg. Banilad. B262). Hildebrandt (L 0108596. Hanauma Bay Beach Park. anon. Barrow Island. 18 March 1992. Villeneuve (SUVA-A 1737).Pagodpud. 25 Feb. J.. leg. N’Yeurt (SUVA-A 5437L).P. 5 June 1967. see text). CG 77b). Cordero and Tuangpalan (US 091395). Dumaguete.Dumaguete City. A. Ti Tree. leg. Amansia rhodantha: n. Agardh). C. Oahu. P.Ilocos Norte. R127). Hakula. – Kenya: Mombassa. 30 Jun. 1996: 472. J. Bay of Sanggar. 16 March 1991.Revision of some Amansia and Melanamansia species 233 Table I. J. C. leg.d. leg. – Fiji: ‘Feejee’. Masuda et al. 347). leg.. 9 Sept. 1838–42.23 May 1978.Siaton. Mauritius. Negros Oriental. paloloensis and Melanamansia glomerata specimens examined in this study. J. 1838–42. leg. Figs 4–10 . 1997. leg. E.7 54.7 2 2 1 2 2 2 2 2 2 * As measured on an index of (1 = little) to (3 = pronounced) – Rotuma: Hoféa. Nanuyalevu. leg. Tantabiddi. 3 Feb. Naukathura Island. leg. Magnesia Virae. Kauai. leg. leg. 1981.. LD 42600 / 0772). JH240). Cannon (SAP 052062). figs 37–42. 2000: 188.Canipo Is.Cuyo. 21/22-Sept. SIPHILEXP-78 (US 088530). 3 and 15 Oct. leg. H. leg. B. Isaac (L 0108594). Plongeurs IRD (AL 288a. leg. 1995. Yasawa Group.Calindagan Reef. 1979.. n. in TCD (Telfair. E. Garrigue (CG 77c. W. leg. Jurien Bay WA. 6 Jun 1978. leg.d. 07 Oct. Average values of selected characters for Amansia paloloensis.6 3. c.d. – New Caledonia: Lajeu Sudouest. 84. Candelaria Zambales. G. G. Isaac (L 0108595). 84 (L 0108603). 1876. 1994 (SUVA-A 797). 1954. Inhaca Peninsula. 24 Nov. G. Suva Reef. SIPHILEXP-78 (US 088534).14 May 1979. Cordero and Tuangpalan (US 091394). Rukuruku Village. 1979. 1834). Montebello Islands. SIPHILEXP-79 (US 088533). leg. leg. J. B. 1980. 1982. leg. Jul. NR 248). 1876. Norris 1988b: 211. Lawai Kai. – New Caledonia: Lajeu Sudouest. leg. C. S. H. 10 Aug. Oahu.Malo. Wilkes Exploring Expedition (US 04057a). 1984. Agardh 1841: 26. South (SUVA-A 2460). A. D. Amansia rhodantha and Melanamansia glomerata surveyed in this study. (US 02890). 1067). W. leg. leg. leg. B79). R. R. Catanduanes. Hildebrandt (L 0108600). Negros Oriental. Carlson (SUVA-A 281). Keats. 27 Dec.Negros Oriental. Ningaloo.leg. 1985. P. L 0108598). leg. Villeneuve (SUVA-A 1774). Garrigue (CH1). Huisman (MURU. Kasahara (SUVA-A 1013. 84. M. Basay. 1984. 1965. Hapmak. leg.leg. Oahu.11 Feb.9 18. J. leg. Viti Levu.Moreland (US 50851). 10 July 1992. Potipot Is. n. 3 Jun 1978. leg. Magruder (US 086459). leg. 28 Nov. AL 288b). Littler (US 163490). – Fiji: ‘Feejee’. 29 May 1978.. 1995. A. Species Thallus height (cm) Blade width (mm) Blade length (mm) Serrations (1 –3)* Cortication (1 –3)* Rosettes (1 –3)* Amansia rhodantha Amansia paloloensis Melanamansia glomerata 39. fide G. Ballou (SUVA-A 5438L). 16 Oct. N’Yeurt (SUVA-A 346. W. Suva Barrier Reef. – Mauritius: Delesseria rhodantha Harvey (holotype) (= Amansia rhodantha): Cap Malheureux. R. 25 September 1999. South (SUVA-A 5431).5 3.d. Yasawa Islands..). A.d. South (SUVA-A 5544.Siquijor Is. SIPHILEXP-78 (US 088532). R.1 22. leg. 1980.Palawan. Farlow (L 0108597). n. Taxonomic results Amansia rhodantha (Harvey) J. – Philippines: Actin. 16 Nov. leg. leg. Cribb (SAP 051714). Sumbawa. Vatualailai. Wilkes Exploring Expedition (US 04057b) (doubtful record. M de Robillard (L 0108599). 14 March 1998..W.Solong-On. Kasahara (SUVA-A 1012. P. 1983. 13 July 1972. Bahati. 06 Sept. figs 1–11 (as Amansia glomerata C. A. A. 1988. E. (MURU. leg. Lopta. 26 Nov. leg. Tangangge. leg. 14 Feb. Kauai. W. – Mozambique: Ponta Abril. leg. 24 April 1996. H. 1982. 17 Jul. 1985. 1995. M. leg. SIPHILEXP-78 (US 013760). Snellius-II (L 0108601). R.. Huisman (CWC 1997. Viti Levu. Daida (US 086755). J. leg. 1973. 10 May 1978. Gaudichaud (‘Amansia glomerata’. Diani Beach. – Philippines:Gao-oa. leg. Huisman. Richer de Forges (AL 288d-CG 77a). 1069). Skelton and G. G. – Kenya: Mombassa. 3 and 5 March 1992. 15 Oct. Garrigue (SUVAA 5436L). Barrow Island.9 11. Chesterfield. Table I presents the average results of morphological measurements carried out on Amansia rhodantha. leg. C. – Indonesia: Cape Batu Kerapo. Jul. leg. Ovalau. 17 Mar. Beqa Lagoon. SIPHILEXP-78 (US 088531). Salayer. Eagles Nest.2 53. leg. leg.

6.5 mm). . US 02890 (scale bars = 3 mm). Habit of mature plant from the type locality. Mauritius. Fig. 5. Detail of tetrasporangial stichidia on lateral endogenous branch. Amansia rhodantha. showing both lax and dense growth forms. from Rotuma Island. N’Yeurt Figs 4–10. D. 8. showing marginal serrations. in TCD (scale bar = 200 µm). Secondary branchlet. in TCD (scale bar = 25 µm). Fig. 9. Rehydrated type material of Delesseria rhodantha Harvey. Indian Ocean. in TCD (scale bar = 2. 10. Fig. Type material of Delesseria rhodantha Harvey. Fig. SUVA-A 5437L. Type material of Delesseria rhodantha Harvey. Surface view of segment cells of blade alae. Figs 4.234 A. showing axial cell (a) surrounded by five pericentral cells. R. Axial structure of blade. and a pair of budding branchlets from the midrib in the lower half. 7. South Pacific (scale bar = 10 µm).

Habit of mature plant. with numerous rosette axes. Fig. Detail of blades. Tetrasporangial stichidia of Amansia rhodantha from Beqa. SUVA-A 1565 (scale bar = 25 µm). Fiji. Fig. SUVA-A 3892 (scale bar = 2.5 mm). Note marginal cystocarps (arrowheads). 4. Segment cells of blade alae in surface view. . Detail of morphologically distinct mature rosette and corticated axis. 12.Revision of some Amansia and Melanamansia species 235 Figs 11–16. Compare with Fig. Amansia paloloensis from Samoa. 14. Detail of the holotype (SUVA-A 1565). Fig. 15. showing both lax branching and the presence of rosettes (arrowheads) (scale bar = 4 mm). Fig. 13. showing lateral endogenous branches and midrib. Fig. SUVA-A 5479L (scale bar = 1 mm). SUVA-A 3892 (scale bar = 12 mm). 16. Holotype of Amansia paloloensis. 11. SUVA-A 5436L (scale bar = 50 µm). Fig.

Fig. fig. CG77a (scale bar = 1 mm). Tetrasporangial stichidia are pinnately arranged on either side of blade margins. 1995: 150. 128. 1. Norris 1995: 66) is usually 4. with more or less pronounced marginal serrations 1–4 mm long (which represent endogenous branches) always present. Fig. containing up to 10 Figs 17–24. 1987: 61. fig. figs 2–14 (type locality: Palolo Deep Marine Reserve. pl. The central axial cell is 28–30 µm in diameter. becoming narrower and disappearing towards the apex. Fig. Synonymy: Rytiphlaea rhodantha (Harvey) Decaisne 1842: 358. denuded below. US 091395 (scale bar = 5 mm). Procarps and curved spermatangial heads are 200–450 µm in diameter and are dorsal on the involucral tips of endogenous lateral branches. and two displaced rows of regularly arranged hexagonal alar cells 20–29 × 92–129 µm in surface view. Fritsch 1945: 570. 24. Cribb 1983: 106. The main axis is irregularly branched. 6. in up to 10 regularly disposed pairs. XXV. showing smooth margins and elongate shape. 85–110 µm thick. 25–42 × 83–125 µm. Individual tetrasporangia are spherical and tetrahedrally divided. 31 fig. N’Yeurt 1996: 428. 100–112 µm in diameter. Isaac 1956: 188. 197. Payri and Meinesz 1985: 511. 1996: 87. New Caledonia. Known distribution: Hawaiian Islands. Plants are brownish-red.5–0. Kenya. Axial structure of plant from Lajeu. – Melanamansia glomerata (C. Falkenberg 1901: 416. 19. New Caledonia. 9). pl. Millar et al. figs 19–21. Melanamansia glomerata. Secondary blades develop as successive pairs on the dorsal side of the midrib. 436B. figs 1–13. South-East Asia. Apia.8 mm in diameter. 21. Fig. fig. A central midrib is present. Blade margins are smooth. Payri and N’Yeurt 1997: 897. . and are composed of lanceolate to ovate blades to 100 mm long and 3 mm wide. Womersley and Bailey 1970: 336. Vietnam. Blades of plant assumed to be from the Fiji Islands (see text). Note that the stichidia is twice laterally branched. 185. 118C–J. Agardh 1824: 194. and the apex is characteristically in-rolled. The number of segments between alternating veins (determinate branches covered by the corticating pericentral cell derivatives. fig. N’Yeurt et al. US 4057b (scale bar = 1 mm). Isaac and Isaac 1968: 24. Abbott 1999: 404. 1. colourless and uniseriate dorsal trichoblasts up to 8 cells long develop on every second or third axial cell (or on every segment in determinate marginal branchlets). pl. AL288b (scale bar = 5 mm). New Caledonia. Philippines. Detail of blade of plant from Mombassa. Coppejans and Millar 2000: 333 (unverified record). 18. and are crisp and lanceolate. figs 127. The number of segments between alternating veins (determinate branches covered by the corticating pericentral cell derivatives. Fig. 267.5 µm in diameter that lies adjacent to the axial cell (Figs 3. pl. 0. R. 4B. surrounded by five pericentral cells. Yamada and Tanaka 1938: 86. Dublin). holotype in TCD. East Africa. Ultimate branchlets are ecorticate. 23. figs 1–3 (type locality: Cap Malheureux. Kylin 1956: 544. Fig. Plant from Potipot Island. Jaasund 1976: 131. A distinct midrib up to 1 mm wide is present in the lower half of ultimate branchlets. Kenya. Yoshida et al. Fig. Fig. fig. E. Tetrasporangia occur in flattened. Plant from Mombassa. with the first two dorsal pericentral cells each developing a smaller pseudo-pericentral cell 13. Okamura 1900: 71. 77–110 µm thick. Agardh) R. Norris 1988b: 211. fig. Millar 1999: 523 (unverified record). New Caledonia. up to 750 µm long and 250 µm broad. 20. Amansia paloloensis South et Skelton 1999: 247. with in-rolled leaf apex.8–14. Internal structure is cellular. 22. up to 100 (on average 40) mm high. 7 mm broad. 17. Japan. showing pseudo-pericentral cells (arrowheads) surrounding (but not directly connected to) axial cell (a). D. Tropical Pacific Ocean. 198. Kenya. Carposporangia are lanceolate. 20. composed of two overlapping layers of elongate hexagonal cells in transverse rows. pl. L 0108598 (scale bar = 5 mm).4. plate CXXVI. Silva et al. 19). Lewis and Norris 1987: 22. e–i. Garrigue and Tsuda 1988: 63. Tetrasporangial stichidia of plant from Oahu. E. The thallus is rose-red. 212a. with a range of 1–5. Figs 11–16 Misapplied names: Amansia glomerata C. Philippines. pinnately arranged marginal stichidia up to 340 µm long and 290 µm in diameter. up to 35 mm long and 6 mm broad. Plant from Unia. Melanamansia glomerata (C. or with regular serrations (endogenous branches). CG77a (scale bar = 10 µm). with a central axial cell 28–30 µm in diameter surrounded by five pericentral cells 36–40 µm in diameter (Figs 1. Norris 1995: 67. while cystocarps 700–1000 µm in diameter occur usually singly and terminally on endogenous branches. figs 1–11. stem-like and rigid in basal portions. Fiji. Weber-van Bosse 1923: 369. and are initially vesiculate. N’Yeurt Basionym: Delesseria rhodantha Harvey 1834: 151–152. Deciduous. Agardh) R. Distribution: Tropical Indian Ocean. L 0108598 (scale bar = 700 µm). figs 14–29. Trichoblasts are sparse. each cell is about 20–20 × 87–100 µm in surface view. with a range of 3–6. SUVA-A 5431 (scale bar = 50 µm). Norris 1995: 67. Tsuda and Wray 1977: 103. Blade of plant from Lajeu. Samoa). Mauritius. 6E (unverified record). Norris 1995: 66) is on average 4. on average 53 mm high. Agardh 1824: 247. Branching is lax to profuse and usually forms deep-red leafy rosettes of secondary branchlets when fully mature. they are up to 4 cells long and non-vesiculate. to 0. The stem of plants is thick and cartilaginous. Marginal teeth are up to 1 mm long and 445 mm broad. 1999: 573.236 A. 2. Figs 17–24 Basionym: Amansia glomerata C.

Revision of some Amansia and Melanamansia species 237 pairs of tetrahedrally-divided tetrasporangia which are 50–100 µm in diameter. procarps and cystocarps occur on endogenous marginal branches. and stained the herbarium mounting paper. the colour of the Melanamansia plants was noticeably brownish-red. Spermatangial heads are curved and arranged in marginal clusters. In all cases. .

it appears to be conspecific with the latter species and thus extends the status of A. fig. figs 1–4) indicate features quite similar to those found in the Amansia species considered in this study from localities in both the Indian Ocean and the tropical Pacific Ocean. Rotuma. the absence of blades organized in morphologically distinct tight axes around the parent axis. Fig. as well as Amansia species from Taiwan. However. Characters such as the degree of rosette formation. a comparison of material of A. The Wilkes collection consists of two plants. degree of cortication of the midrib and position of the reproductive stichidia are equally quite variable in plants from the same and similar populations. Smith (1979: 38–40) points out that the accounting and . with for instance A. an additional field test to distinguish Amansia spp. while the other (US 04057b) is Melanamansia glomerata. and absence of pseudopericentral cells. Mozambique and Western Australia. a single record for this genus exists (collected from ‘Feejee’ by the Charles Wilkes U. having tight rosettes as well as very lax. and a cross section of the blade clearly shows the five pericentral cells around the axial ring. 14) and corticated rosette axes (Fig. rhodantha. The status of Amansia paloloensis South et Skelton South and Skelton (1999) described a new species of Amansia from Samoa. com.S. In the Fiji Islands. rhodantha from the Indian Ocean and other localities. These plants also had blades with well-developed midribs in the lower half (Fig. Now that the Samoan species has been compared with genuine herbarium material of A. Eastern Australia. paloloensis. length of the blades. generally smaller. paloloensis showed no major difference in the range of characteristics among them. and trichoblasts lacking protective vesicles when young (Norris 1988b). Agardh from Hawaii (LD 42600 / 0772) clearly shows the presence of paired dorsal pseudo-pericentral cells in a blade cross section (Fig. and a weakly developed midrib as characters distinguishing A. more coriaceous branchlets with less apparent midribs (this study). Vidalia. The present author has re-examined the holotype. rhodantha from Western Australia (MURU B262. Melanamansia glomerata in the tropical Pacific Wilson and Kraft (2000: 366) commented that the generic segregation of Melanamansia from Amansia is probably not warranted. Exploring Expedition. Kenya. while Melanamansia becomes dark brown. additional distinguishing characteristics of Melanamansia are a darkly coloured brown thallus which stains the mounting paper when dried (possibly linked to differences in chemical composition. loriform blades reminiscent of some specimens of A. On the basis of these observations. More importantly. and these authors considered the lack of highly corticated axes. paloloensis. from Melanamansia glomerata was found: when preserved in 4% Formalin-seawater and exposed to ambient light for several days. material from the Tropical Pacific and Indian Oceans previously known as Amansia glomerata (or misidentified as Melanamansia glomerata.238 A. rhodantha (South and Skelton 1999: 249). R. a re-examination of the lectotype of Amansia glomerata C. 12). the Philippines. and Ceramium. between 1838 and 1842 (US 04057b. Fiji. While there is no doubt as to the generic identity of the latter specimen. Japan. thicker. The reproductive and trichoblast anatomy of A. Masuda pers. MURU 248). D. paloloensis from A. plate CXXVI. indicating that the morphology of the species is influenced perhaps by environmental factors. rhodantha from the type collection and localities in Mauritius. From this study. Amansia tends to bleach to translucent white. New Caledonia.13). rhodantha as described and illustrated by Norris (1988b: 211). when pseudo-pericentral cells were found to be absent) are considered conspecific with Amansia rhodantha. serrate branches and lax ‘rosettes’ of secondary blades issued from the weakly developed midrib. 3) which distinguishes that plant from A. since the presence of pseudo-pericentral cells is an inconsistent attribute in species of Neurymenia. 2) indeed shows a plant with loriform. M. 23) but the plant has not been collected in Fiji waters since. The drawing of the holotype (South and Skelton 1999: 246. N’Yeurt Discussion The holotype of Delesseria rhodantha Harvey An examination by the author of a rehydrated blade fragment of the holotype of Delesseria rhodantha Harvey in TCD (Fig. isotype and other collections of A. paloloensis from the type locality. rhodantha as a pan-tropical entity. On the other hand. The habit and description of the holotype by Harvey (1834: 151. regularly alternate marginal serrations which are not as pronounced as in Amansia rhodantha and which are often absent. Kuetzingia. one of which (US 04057a) is Amansia rhodantha. including the presence of both tight and lax rosettes of secondary branches and alar cells on the primary axes. is comparable to A. 7) shows features similar to those seen in blades of South Pacific material previously ascribed to Amansia glomerata. apparently because the brown pigments (presumably polyphenols) are less susceptible to the bleaching action of the preservative (as is the case in species of brown algae such as Sargassum and Turbinaria). which is welldocumented and illustrated in the protologue of the species. the Cook Islands and French Polynesia (Table I) with A. and found a significant number of thalli with morphologically distinct primary corticated axes and relatively tight and lax rosettes of secondary blades on rosette axes (Figs 11.).

Validity of current criteria to distinguish the Amansieae Characters that have proved useful in distinguishing species in the Amansieae include the presence or absence of pseudo-pericentral cells. type of endogenous branching. older plants tend to have thicker. except the Hawaiian samples. and a main axis with alar cells not very distinct from secondary branches. the description of the species states that branches have the same shape as the main axis (i. and found to lack pseudo-pericentral cells. These ventral. L 0108603). and found to consistently lack pseudo-pericentral cells and a similar observation was made on samples from Taiwan. in the case of A. rhodantha. which was the main reason for separating the species from A. 5. paloloensis revealed a wide range in the degree of rosette formations and cortication of the secondary axes and presence of alae on primary axes. fig. 2). and would seem to function as attachment haptera. Indian Ocean Amansia rhodantha also lacked such cells. 1) shows both the presence of tight and lax rosettes of secondary branches. thus placing them in Amansia. in New Caledonia and Kenya). when present). The author has unsuccessfully tried to obtain material from the latter locality. irregularly dichotomous proliferations lack alar cells. Indeed. In most instances. which also had a consistently thicker thallus. pseudo-pericentral cells were universally absent. glomerata occurs in the same habitat as Amansia rhodantha (e. Payri and N’Yeurt 1997) were reexamined. 7). Melanamansia glomerata is unlikely to be confused with other Melanamansia species because of its unique combination of characteristics (Norris 1988b. and the presence of tight and lax rosettes of secondary branches on the same plant (Figs 11–13). and is backed by other morphological and biochemical differences (Norris 1988b. with the blades smooth and superficially reminiscent of those of some Sargassum species (Figs 22–24). Indonesian and Indian Ocean region were sectioned and examined for pseudo-pericentral cells. so it cannot be confirmed at this stage if M. and in some instances lacking altogether. calm waters (such as blue holes. 41) similarly report the absence of pseudo-pericentral cells in A. fig. they were clearly apparent and about half the size of pericentral cells (Fig. which is a characteristic feature of both Amansia and Melanamansia (Norris 1988b) can vary from lax to tight. However. Masuda. com. (2000: 188. Ecological studies would be needed to verify these hypotheses. 1995). rhodantha from Mauritius and Western Australia (compare Figs 4. in the case of Amansia paloloensis). Herbarium and liquid-preserved samples of Amansia and Melanamansia species from the Pacific. All Amansia rhodantha specimens examined had fairly prominent. more corticated secondary and primary axes with tight rosettes.). some specimens were virtually indistinguishable from specimens of A. CXXVI. It would seem that perhaps shallow. both on the same plant and within individuals in similar populations. Proliferations from the ventral midrib (aside from secondary branchlets) were only found to be abundant in some of the Indonesian Amansia rhodantha specimens from Salayer (e. When pseudo-pericentral cells were present. in some cases. 19). 1996. In Melanamansia glomerata these marginal serrations were much less pronounced (but more regular than in Amansia rhodantha. while in Malaysia Masuda et al. adding to the confusion between the two genera. The presence or absence of these pseudo-pericentral cells hence appears to be a good generic criterion. Herbarium specimens of ‘Amansia glomerata’ from southern Japan housed in SAP (Hokkaido University) were examined. alternate marginal proliferations (which represent non-determinate or endogenous secondary branchlets) 2–5 mm long (Fig. environmental factors seem to play a role in determining the degree of rosette formation. pers.g. pl. glomerata occurs in the Solomon Islands. Fiji and New Caledonia (Table I). However. position of the reproductive axes and the distinctness of the axial ring in rosette axes (Norris 1988a. 13). perhaps an adaptation to the exposed habitats where the plants came from.g. the Philippines. paloloensis. degree of marginal serrations and thallus flattening. For instance. Also. however it is clear that the degree of rosette formation and cortication of the axes is not a reliable taxonomic character at the species level in Amansia. From an examination of the specimens at hand. and some samples from Kenya. Norris (1988b: 211) reported that rosette axes in . the vesiculate (in Amansia) or non-vesiculate (in Melanamansia) nature of trichoblasts. M. and hence the precise locality and date of this Melanamansia collection from Fiji is unknown.e. it was observed that rosettes of blades along the secondary axes. the presence or absence of bladelets in rosettes. There is a possibility it might have been mixed with collections from neighbouring localities such as the Solomon Islands. rhodantha. an examination of the holotype and liquid-preserved and pressed isotypes of A. fig. Fijian and French Polynesian specimens previously ascribed to Melanamansia (N’Yeurt et al. while younger plants have less cortication and more lax or absent rosettes of blades.Revision of some Amansia and Melanamansia species 239 recording of the Wilkes Expedition collections was not very satisfactory. more exposed conditions (such as on fringing reefs) favour tight rosette formation (which is more hydrodynamically resistant) while more lax arrangements are possible in deeper. M. The illustration by Harvey of the holotype of Delesseria rhodantha (1834. Generally. Wilson and Kraft 2000). both have an ala) and are arranged in ‘lax’ rosettes (South and Skelton 1999: 247.

This feature was examined on a range of specimens of Amansia from the South Pacific and the Indian Ocean but no significant difference was seen in these specimens. rhodantha and A. only being more or less surrounded by cortication. 16) and Melanamansia show no particular difference. From Table I. Biogeographical map for Amansia rhodantha (circle) and Melanamansia glomerata (triangle). These differences reinforce the conspecificity of the Amansia rhodantha and A. There appears to be no notable difference in the range of thallus size.).4 (Norris 1995). as well as between A. Norris (1995: 66) counted the number of segment cells between these alternating veins for several species of Melanamansia and Amansia. as the axial ring was still discernible in all sections made. rhodantha specimens observed in this study. rhodantha studied. smaller and narrower blades. cortication of axes and rosette formation between specimens of Amansia rhodantha studied. blade dimensions. as well as more corticated axes. and found them relatively constant. showing the currently known distribution. but can also occur apically. However. An examination of these elegant cells in species of Amansia (Figs 10. At this stage there seems to be no largely significant difference in the number of segments between alternating veins in the specimens of A. R. Fig. but the range is reportedly less than in Amansia rhodantha. . in elongate flattened and pinnate stichidia which are usually on lateral marginal branchlets. paloloensis specimens studied. 15). com. with most specimens having segment lengths of about 100 µm.240 A. in that they are typically rhodomelaceous with pairs of tetrasporangia in up to ten rows. 20). Tetrasporangial stichidia are basically similar in both Amansia rhodantha and Melanamansia glomerata. paloloensis. a number of preliminary conclusions can be made: A. N’Yeurt Amansia from Natal have an ill-defined axial row of cells. D. as well as the generic distinctness of Melanamansia glomerata. which was not seen in Amansia rhodantha (Figs 6. An additional potentially useful character is the number of segments between lateral ‘veins’ on the blades. it was noted that in Melanamansia the range was more restricted. Hence the distinctness of the axial ring was not found a reliable taxonomic character among the species examined. B. These can be easily detected in the unrolled apical portions of the blades by the developing trichoblast terminating each such alternating ‘vein’ (Norris 1995 and pers. In Melanamansia it averages 4. In A. with the length of segments ranging from 80 to 120 µm. the number ranged from 3 to 7.3. Melanamansia glomerata from all localities studied have consistently smoother. This also varied according to the degree of rosette formation and age of the plant. which was also recorded by Norris (1988b) for Hawaiian material. which are in fact determinate branches covered by the pericentral cell derivatives. 25. Cover cells usually totally cover the developing sporangia. and occur in two overlapping and slightly offset layers radiating on either side of the midrib. usually 3 or 4 with an average of 3. One feature which was noted in Melanamansia glomerata is that individual tetrasporangial stichidia are often laterally branched once or twice (Fig. Cells of the blade alae in both Amansia and Melanamansia are regularly elongate-hexagonal.

M. Foreword. Die Rhodomelaceen des Golfes von Neapel und der angrenzenden Meeres-Abschnitte. J. Fauna und Flora des Golfes von Neapel. 1983. South pers. 1968. Lund).Vietnam and Malaysia (M. U. Lewis. Myxophyceae. J. University of Tromsø. Agardh. from “Cap Malheureux”. A. Scheltema & Holkema. P. and F.W. East Africa Nat. rhodantha and Melanamansia glomerata are more widely distributed than previously thought. K.. Masuda. and R. S. S. Sci. Schofield. and Dr A. Tsuda.. K. Bishop Museum Press. Paris 1: 330–333. Annal. 2000. E. N. Hist. Die Gattungen der Rhodophyceen. M. E. Australian Coral Reef Society: Brisbane. com. J. J. ser. Robin South is also thanked for his helpful comments on an earlier draft of this manuscript. Phang. Shimada. Cambridge University Press. Bot. Professor Willem Prud’homme van Reine (L – Rijksherbarium. Holmgren. sparse. xvi + 477 + [1] pp.). C. 1981. 1956. composed of up to 4 cells: Melanamansia glomerata. Phil. W. II. Volume II. Nat. (Hooker) 1: 147–157. Mr Barrett Brooks and Dr John Sims (US – Smithsonian Institution. Bot. Charles Telfair. et description de cinq nouveaux genres de cette famille. E. 336 figs. Rhodophyta.). C. The Structure and Reproduction of the Algae. 3. Decaisne. Marine Algae of the Southern Great Barrier Reef. Acknowledgements The author sincerely thanks the following persons and institutions for the generous loan of type material and specimens used in this study: Dr John Huisman (MURU – Murdoch University). 71 pls. Part I. com. 1945. Micronesica 21: 53–70. F. W. pls v–vii. 10 figs. Utrecht. Revised Key to the Species of Amansia and Melanamansia in the South Pacific – Presence of paired dorsal pericentral cells. Dr Claude Payri (UPF – Tahiti). First edition. Washington). Marine algae of Inhaca Island and of the Inhaca Peninsula. Kato. xvi + 754 pp.Revision of some Amansia and Melanamansia species 241 Conclusions The results of this study and those of South and Skelton (1999) and of Masuda et al. J. T. in the Mauritius. 22: 161–193. Phaeophyceae. Nouv. 2 folded maps. 1809. iii + 159 pp. V. 1824. E. J. It is clear that both A. Ms Susanna Riebe (LD – Botanical Museum. New Caledonia and the Philippines (this study). Part 1. Pub. A. Nat. C. E. 17: 297–380. Rhodophyceae. M. B. Kenya (this study). K. Coppejans. K. Hommersand. 2000. J. Berlin. Falkenberg. Norris is thanked for his helpful insights and suggestions for this research. Lord Howe Island (G. E. and possibly other localities which are not yet adequately investigated. Taxonomic notes on marine algae from Malaysia. Contrib. Bohn. Jaasund. Kylin. Bull. flora and vegetation. pig- References Abbott. 443–457. Mar. Sapporo). Figure 25 presents a world distribution of these two genera as it is currently known. 1841. Norris. Sci. Mar. 29: 1–38. Index Herbariorum. 1988. Isaac. Cal. 1901. A history and annotated account of the benthic marine algae of Taiwan. Isaac. xiv + 939 pp. Smiths.. Notice of a collection of algae. with descriptions of some new and little known species. . Essais sur une classification des algues et des polypiers calcifères de Lamouroux. Observations sur la physiologie des algues marines. 1842. 1999. Keuken and E. Systema algarum. trichoblasts non-vesiculate. African Bot. initially vesiculate and composed of up to 8 cells: Amansia rhodantha. – Absence of paired dorsal pericentral cells. Bot. K. 35: 165–366. G. and A. frontispiece. Mus. and each genus is consistently differentiated from the other by a number of anatomical and biochemical characters. H. Garrigue. Kawaguchi and S. Mar. J. H.. E. Millar. Honolulu. Handbook No 2. M. cxxvi.. Linnaea 15: 1–50. In historiam algarum symbolae. J. 1834. 43: 315–346. Cribb. pigments reddish-brown. H. 27: 7–28. Dublin). Sci. Professor G. 458 figs. and J. Intertidal Seaweeds in Tanzania. Marine Red Algae of the Hawaiian Islands. xv + 673 pp. S. Fritsch. Lund. Bot. P. Isaac. Soc. Professor G. Marine red algae from the North coast of Papua New Guinea. Monographie 26. Robin South and Mr Posa A. and it appears that Melanamansia glomerata is so far restricted to Hawaii (Norris 1995). Lamouroux. communicated to Dr. Agardh. 173 + (2) pp. W. Accepted 17 January 2002. 2. 1976. The Herbaria of the World. Dr John Parnell (TCD – Trinity College. (2000) confirm that the genus Amansia does exist in the Pacific Ocean. 43: 181–190.A. pls cxxv. Harvey. Seven species of Rhodophyceae. 1987. W. Marine botany of the Kenya coast.A. I. xxxviii + 312 pp. Soc. Millar for his generous assistance in obtaining copies of rare publications. Catalogue of marine benthic algae from New Caledonia. Lunda (Lund). Dr R. I. Professors Michio Masuda and Tadao Yoshida (SAP – Hokkaido University. General account of the environment. F.. ments rose-red. 7th edition. Hooker by the late Mrs. trichoblasts on every second or third axial cell.. Masuda pers. 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